Time – size tradeoffs: a phylogenetic comparative study of flowering time, plant height and seed mass in a north-temperate flora

Parents face a timing problem as to when they should begin devoting resources from their own growth and survival to mating and offspring development. Seed mass and number, as well as maternal survival via plant size, are dependent on time for development. The time available in the favorable season will also affect the size of the developing juveniles and their survival through the unfavorable season. Flowering time may thus represent the outcome of such a time partitioning problem. We analyzed correlations between flowering onset time, seed mass, and plant height in a north-temperate flora, using both cross-species comparisons and phylogenetic comparative methods. Among perennial herbs, flowering onset time was negatively correlated with seed mass (i.e. plants with larger seeds started flowering earlier) while flowering onset time was positively correlated with plant height. Neither of these correlations was found among woody plants. Among annual plants, flowering onset time was positively correlated with seed mass. Cross-species and phylogenetically informed analyses largely agreed, except that flowering onset time was also positively correlated with plant height among annuals in the cross-species analysis. The different signs of the correlations between flowering onset time and seed mass (compar. gee regression coefficient=−7.8) and flowering onset time and plant height (compar. gee regression coefficient=+30.5) for perennial herbs, indicate that the duration of the growth season may underlie a tradeoff between maternal size and offspring size in perennial herbs, and we discuss how the partitioning of the season between parents and offspring may explain the association between early flowering and larger seed mass among these plants.     

Determinants of Dispersal Distance in Free-Ranging Juvenile Lizards

Dispersal of offspring from their natal site has a critical influence on individual fitness. Although the consequences of dispersal have received much theoretical attention, the determinants of dispersal remain poorly understood for many animals. To address this issue, we marked and released size‐manipulated hatchling lizards (Amphibolurus muricatus; Agamidae) over a 3‐mo period in the field to evaluate the effects of body size and the time of hatching on dispersal distance. Our mark–recapture data indicated that body size and offspring sex had little effect on distances travelled by individuals. However, the timing of hatching had a strong impact; individuals that hatched early in the season dispersed further than did those hatching late. This pattern may allow early‐hatched juveniles to disperse and secure high‐quality habitats before the arrival of later‐hatched conspecific competitors.     

Parental effects in Plantago lanceolata L. III. Measuring parental temperature effects in the field

To determine the evolutionary importance of parental environmental effects in natural populations, we must begin to measure the magnitude of these effects in the field. For this reason, we conducted a combined growth chamber-field experiment to measure parental temperature effects in Plantago lanceolata. We grew in the field offspring of controlled crosses of chamber-grown parents subjected to six temperature treatments. Each treatment was characterized by a unique combination of maternal prezygotic (prior to fertilization), paternal prezygotic, and postzygotic (during fertilization and seed set) temperatures. Offspring were followed for three years to measure the effects of treatment on several life-history traits and population growth rate, our estimate of fitness. Parental treatment influenced germination, growth, and reproduction of newborns, but not survival or reproduction of offspring at least one year old. High postzygotic temperature significantly increased germination and leaf area at 17 weeks by approximately 35% and 2%, respectively. Probability of flowering and spike production in the newborn age class showed significant parental genotype x parental treatment interactions. High postzygotic temperature increased offspring fitness by approximately 50%. The strongest contributors to fitness were germination and probability of flowering and spike production of newborns. A comparison of our data with previously collected data for chambergrown offspring shows that the influence of parental environment on offspring phenotype is weaker but still biologically meaningful in the field. The results provide evidence that parental environment influences offspring fitness in natural populations of P. lanceolata and does so by affecting the life-history traits most strongly contributing to fitness. The data suggest that from the perspective of offspring fitness, natural selection favors parents that flower later in the flowering season in the North Carolina Piedmont when it is warmer. Genotypic-specific differences in response of offspring reproductive traits to parental environment suggest that parental environmental effects can influence the rate of evolutionary change in P. lanceolata.     

An empirical estimate of the generation time of mouse lemurs

The generation time of organisms drives the rate of change in populations and across evolutionary times. In long‐lived species, generation time should also account for overlapping generations, and the average age of parents has been proposed as a best approximation under these conditions. This study uses this definition to estimate the generation time of a widely studied small primate, Microcebus murinus, based on parentage data generated for a free‐living population over a 6‐year period in northwestern Madagascar. The average age of parents was calculated separately for mothers and fathers of three different offspring cohorts that differed in the degree of demographic uncertainty. In addition, adult survival rates were calculated for males and females based on long‐term capture data from the same population to estimate the possible upper limits of generation time. Adult survival was low with only 44% of adult females and 38% of adult males being recaptured at the beginning of their second breeding season. The average age of mothers was 1.56–1.91 years, pointing toward a 2‐year female generation time due to the high proportion of 1‐year old mothers in all three cohorts. Female generation time estimates were fairly stable across the three offspring cohorts. In contrast, the average age of fathers differed by more than 1 year from the first to the third offspring cohort (1.71–2.83 years) pointing toward a 3‐year generation time, but also suggesting a higher degree of demographic uncertainty in the early years of the study. For future modeling purposes, we, therefore, propose to use the average, 2.5 years, of male and female values as new estimate for the generation time of mouse lemurs.     

GENE FLOW IN CHAMAECRISTA FASCICULATA (LEGUMINOSAE) I. GENE DISPERSAL

Both pollen and seed dispersal components of gene flow were examined in the annual plant Chamaecrista fasciculata (Leguminosae) and quantified in terms of Wright's neighborhood area. Pollen dispersal was estimated by measuring pollinator flight movement throughout the flowering season and the contribution of pollen carryover to pollen dispersal was determined by comparing pollinator flight movement with dispersal of electrophoretic markers in an experimental transect. Phenological effects on the probability of fruit set were measured to determine whether pollinations should be weighted differentially across the flowering season. The outcrossing rate, a major determinant of the role of pollen dispersal in gene flow, was estimated from electrophoretic analysis of progeny arrays and by measuring the proportion of nongeitonogamous pollinator flight movements. Seed dispersal was measured in a prairie habitat and in experimental plots without surrounding vegetation. Seed dispersal was small in comparison to pollen dispersal in both environments. Fruit set was low at the beginning and end of the flowering season, periods when flower density is low and pollinator flight distances are large. Although the outcrossing rate was high (t = 80%) and pollen carryover substantial, pollen dispersal was limited. Averaged over 4 years, neighborhood area, based on both seed and pollen dispersal, was 17.6 m², and corresponds to a circle of radius 2.4 m. The observed limited gene dispersal suggests the population of C. fasciculata is genetically subdivided into small breeding units of related individuals.     

Grassland plant functional groups exhibit distinct time-lags in response to historical landscape change

Recent studies have shown significant impacts of past landscapes on present distributions of species, and discussed the existence of an extinction debt. Understanding of the processes building an extinction debt is fundamentally important for explaining present and future diversity patterns of species in fragmented landscapes. Few empirical studies, however, have examined the responses of different plant functional groups (PFGs) to historical landscape changes. We aimed to reveal PFG-based differences in species’ persistence by focusing on their vegetative, reproductive, and dispersal traits. We examined whether the present distributions of PFGs of grassland species in the edges of remnant woodlands established on former semi-natural grasslands are related to the past surrounding landscapes at different time periods and spatial scales. The effects of past landscapes varied significantly among the PFGs. Richness of short, early flowering forbs and tall, late-flowering, wind-dispersed forbs showed significant positive relationships with the surrounding habitat proportions more than 50 years ago (the 1950s) and at wide spatial scales (more than 1 km²). Richness of tall, late-flowering forbs with unassisted and other types of dispersal mechanisms showed positive relationships with the surrounding habitat proportions in recent times (the 1970s) and at smaller spatial scales (0.25 km²). Our results suggested that plant growth form, flowering season and dispersal ability—with additional information on seed bank persistence—can be good indicators for identifying species’ specific sensitivity to surrounding habitat loss. Trait-based approaches can be useful for understanding present and future distributions of grassland species with different persistence strategies in human-modified landscapes.     

Long-term consequences of early ontogeny in free-living Great Tits Parus major

Environmental factors during early development may have profound effects on subsequent life-history traits in many bird species. In wild birds, sex-specific effects of early ontogeny on natal dispersal and future reproduction are not well understood. The objective of this work was to determine whether hatching date and pre-fledging mass and condition of free-living Great Tits Parus major have any subsequent effect on individuals’ natal dispersal and reproductive performance at first breeding. Both males and females dispersed longer distances in coniferous than in deciduous forests, while dispersal was condition-dependent only in males (heavier as nestlings dispersed farther). In females, mass and condition at pre-fledging stage correlated significantly with clutch size, but not with subsequent reproductive performance as measured by fledging success or offspring quality. In contrast, heavier males as nestlings had higher future fledging success and heavier offspring in their broods compared with those in worse condition as nestlings. The hatching date of female as well as male parents was the only parental parameter related to the number of eggs hatched at first breeding. These results indicate that pre-fledging mass and condition predict subsequent fitness components in this bird species. We suggest that sex-specific relationships between a disperser’s condition and its selectivity with respect to breeding habitat and subsequent performance need to be considered in future models of life-history evolution.     

Reduced mortality selects for family cohesion in a social species

Delayed dispersal is the key to family formation in most kin-societies. Previous explanations for the evolution of families have focused on dispersal constraints. Recently, an alternative explanation was proposed, emphasizing the benefits gained through philopatry. Empirical data have confirmed that parents provide their philopatric offspring with preferential treatment through enhanced access to food and predator protection. Yet it remains unclear to what extent such benefits translate into fitness benefits such as reduced mortality, which ultimately can select for the evolution of families. Here, we demonstrate that philopatric Siberian jay (Perisoreus infaustus) offspring have an odds ratio of being killed by predators 62% lower than offspring that dispersed promptly after independence to join groups of unrelated individuals (20.6% versus 33.3% winter mortality). Predation was the sole cause of mortality, killing 20 out of 73 juveniles fitted with radio tags. The higher survival rate among philopatric offspring was associated with parents providing nepotistic predator protection that was withheld from unrelated group members. Natal philopatry usually involves the suppression of personal reproduction. However, a lower mortality of philopatric offspring can overcome this cost and may thus select for the formation of families and set the scene for cooperative kin-societies.     

Low plant density enhances gene dispersal in the Amazonian understory herb Heliconia acuminata

In theory, conservation genetics predicts that forest fragmentation will reduce gene dispersal, but in practice, genetic and ecological processes are also dependent on other population characteristics. We used Bayesian genetic analyses to characterize parentage and propagule dispersal in Heliconia acuminata L. C. Richard (Heliconiaceae), a common Amazonian understory plant that is pollinated and dispersed by birds. We studied these processes in two continuous forest sites and three 1‐ha fragments in Brazil's Biological Dynamics of Forest Fragments Project. These sites showed variation in the density of H. acuminata. Ten microsatellite markers were used to genotype flowering adults and seedling recruits and to quantify realized pollen and seed dispersal distances, immigration of propagules from outside populations, and reproductive dominance among parents. We tested whether gene dispersal is more dependent on fragmentation or density of reproductive plants. Low plant densities were associated with elevated immigration rates and greater propagule dispersal distances. Reproductive dominance among inside‐plot parents was higher for low‐density than for high‐density populations. Elevated local flower and fruit availability is probably leading to spatially more proximal bird foraging and propagule dispersal in areas with high density of reproductive plants. Nevertheless, genetic diversity, inbreeding coefficients and fine‐scale spatial genetic structure were similar across populations, despite differences in gene dispersal. This result may indicate that the opposing processes of longer dispersal events in low‐density populations vs. higher diversity of contributing parents in high‐density populations balance the resulting genetic outcomes and prevent genetic erosion in small populations and fragments.     

Invader immunology: invasion history alters immune system function in cane toads (Rhinella marina) in tropical Australia

Because an individual's investment into the immune system may modify its dispersal rate, immune function may evolve rapidly in an invader. We collected cane toads (Rhinella marina) from sites spanning their 75‐year invasion history in Australia, bred them, and raised their progeny in standard conditions. Evolved shifts in immune function should manifest as differences in immune responses among the progeny of parents collected in different locations. Parental location did not affect the offspring's cell‐mediated immune response or stress response, but blood from the offspring of invasion‐front toads had more neutrophils, and was more effective at phagocytosis and killing bacteria. These latter measures of immune function are negatively correlated with rate of dispersal in free‐ranging toads. Our results suggest that the invasion of tropical Australia by cane toads has resulted in rapid genetically based compensatory shifts in the aspects of immune responses that are most compromised by the rigours of long‐distance dispersal.     

Behavioral and Social Structure Effects on Seed Dispersal Curves of a Forest‐Interior Bulbul (Pycnonotidae) in a Tropical Evergreen Forest

Movement patterns of animals can vary dramatically as a function of their reproductive cycle or social structure; however, little is known about how changes in the social structure of dispersers affect patterns of seed dispersal. We examined the movement patterns of the forest‐dwelling and cooperatively breeding Puff‐throated Bulbul (Alophoixus pallidus) in relation to different stages of their reproductive cycle, time of day, and group size, to determine potential impacts on the shape and scale of dispersal curves generated using a combination of gut passage time and displacement distance data. There were significant differences in dispersal distances depending on group size, season (breeding, non‐breeding), incubation (vs. other times of the year), and time of the day. The estimated median seed dispersal distance was 28 m. The median dispersal distances produced by birds in larger groups were longer than those of smaller groups (29 m vs. 25 m). During the breeding season, median dispersal distances were longer than during the non ‐ breeding season (31 m vs. 25 m), but the median dispersal distances were significantly shorter during incubation than during outside incubation (24 m vs. 28 m). The median dispersal distance produced in the early morning (30 m) was also longer than that of other times of the day (23 m late morning, 28 m early afternoon, and 26 m late afternoon). This study suggests that various aspects of an animal's behavior are likely to have significant effects on seed shadows and that this may vary significantly even among individuals of the same species.     

Fine-scale genetic structure and gene dispersal in Centaurea corymbosa (Asteraceae) I. Pattern of pollen dispersal

Pollen dispersal was characterized within a population of the narrowly endemic perennial herb, Centaurea corymbosa, using exclusion-based and likelihood-based paternity analyses carried out on microsatellite data. Data were used to fit a model of pollen dispersal and to estimate the rates of pollen flow and mutation/genotyping error, by developing a new method. Selfing was rare (1.6%). Pollen dispersed isotropically around each flowering plant following a leptokurtic distribution, with 50% of mating pairs separated by less than 11 m, but 22% by more than 40 m. Estimates of pollen flow lacked precision (0-25%), partially because mutations and/or genotyping errors (0.03-1%) could also explain the occurrence of offspring without a compatible candidate father. However, the pollen pool that fertilized these offspring was little differentiated from the adults of the population whereas strongly differentiated from the other populations, suggesting that pollen flow rate among populations was low. Our results suggest that pollen dispersal is too extended to allow differentiation by local adaptation within a population. However, among populations, gene flow might be low enough for such processes to occur.     

Parentage analysis detects cryptic precapture dispersal in a philopatric rodent

Locating birthplaces using genetic parentage determination can increase the precision and accuracy with which animal dispersal patterns are established. We re-analyse patterns of movement away from the birthplace as a function of time, sex and population density for a sample of 303 banner-tailed kangaroo rats, Dipodomys spectabilis. We located birth sites using a combination of likelihood-based parentage analysis with live-trapping of mothers during the breeding season. The results demonstrate that natal-breeding site distances are density dependent in this species; in particular, both sexes emigrate earlier in the year, and females disperse farther than males, at low population densities. Banner-tailed kangaroo rats were chosen as a study system because live-trapping easily detects maternal and offspring locations; nevertheless, parentage analysis reveals that some offspring evade early detection and move substantial distances before their first capture. In a few cases, the approach even detects dispersal out of the natal 'deme' prior to first capture. Parentage analysis confirms the extreme philopatry of both sexes but indicates that prior estimates of median dispersal distance were too low. For D. spectabilis, more accurate location of individual birthplaces clarifies patterns of sex bias and density dependence in dispersal, and may resolve apparent discrepancies between direct and indirect estimates of dispersal distance. For species in which mothers can be more reliably trapped than juveniles, using offspring genotypes to locate parents is a novel way that genetic techniques can contribute to the analysis of animal dispersal.     

Seed dispersal and seed banks in Aloe marlothii (Asphodelaceae)

Aloe marlothii flowers during dry winter months (July–September) and produces large numbers of wind dispersed seeds. Fire disturbance in a population of several thousand A. marlothii plants at Suikerbosrand Nature Reserve, Gauteng, permitted a series of seed dispersal experiments to be conducted. Germination trials indicated that seedling emergence decreased with increased distance from a well defined aloe stand and burn area margin, with seeds dispersed up to 25 m. Flowering frequency and total seed production were positively correlated with plant height, with seed production estimated to range from 26,000 to 375,000 seeds/plant. Although a large number of seeds are produced by flowering plants the survival rate of seeds did not extend beyond the following flowering season.     

Maternal effects provide phenotypic adaptation to local environmental conditions

In outcrossing plants, seed dispersal distance is often less than pollen movement. If the scale of environmental heterogeneity within a population is greater than typical seed dispersal distances but less than pollen movement, an individual's environment will be similar to that of its mother but not necessarily its father. Under these conditions, environmental maternal effects may evolve as a source of adaptive plasticity between generations, enhancing offspring fitness in the environment that they are likely to experience. This idea is illustrated using Campanula americana, an herb that grows in understory and light-gap habitats. Estimates of seed dispersal suggest that offspring typically experience the same light environment as their mother. In a field experiment testing the effect of open vs understory maternal light environments, maternal light directly influenced offspring germination rate and season, and indirectly affected germination season by altering maternal flowering time. Results to date indicate that these maternal effects are adaptive; further experimental tests are ongoing. Evaluating maternal environmental effects in an ecological context demonstrates that they may provide phenotypic adaptation to local environmental conditions.     

Parentage assignment detects frequent and large-scale dispersal in water voles

Estimating the rate and scale of dispersal is essential for predicting the dynamics of fragmented populations, yet empirical estimates are typically imprecise and often negatively biased. We maximized detection of dispersal events between small, subdivided populations of water voles (Arvicola terrestris) using a novel method that combined direct capture-mark-recapture with microsatellite genotyping to identify parents and offspring in different populations and hence infer dispersal. We validated the method using individuals known from trapping data to have dispersed between populations. Local populations were linked by high rates of juvenile dispersal but much lower levels of adult dispersal. In the spring breeding population, 19% of females and 33% of males had left their natal population of the previous year. The average interpopulation dispersal distance was 1.8 km (range 0.3-5.2 km). Overall, patterns of dispersal fitted a negative exponential function. Information from genotyping increased the estimated rate and scale of dispersal by three- and twofold, respectively, and hence represents a powerful tool to provide more realistic estimates of dispersal parameters.     

Possible influences of habitat characteristics on the evolution of semelparity and cannibalism in the hump earwigAnechura harmandi

The possible influences of life history and habitat characteristics on the evolution of semelparity and cannibalism in the hump earwigAnechura harmandi were studied. This species is univoltine and overwinters as an adult. Females laid single egg-batches during winter in nests under stones at a riverside in a valley. They took care of the eggs which hatched in early spring and the offspring ate their mother before dispersing. The valley was sometimes flooded in summer. Nymphs emerged as adults and dispersed to elsewhere before the rainy season arrived. They returned to the riverside after the rainy season. The flooding and/or summer heat seemed to be the selective force for the evolution of dispersal behavior and semelparity in this species. The cannibalism of the female parent by her offspring seemed to have readily evolved after the evolution of semelparity. The unfavorable environmental conditions seemed to have a large effect on the evolution of semelparity and cannibalism in this species.     

What drives selection on flowering time? An experimental manipulation of the inherent correlation between genotype and environment

The optimal timing of the seasonal switch from somatic growth to reproduction can depend on an individual's condition at reproduction, the quality of the environment in which it will reproduce, or both. In annual plants, vegetative size (a function of age at flowering) affects resources available for seed production, whereas exposure to mutualists, antagonists, and abiotic stresses in the environment (functions of Julian date of flowering) influences success in converting resources into offspring. The inherent tight correlation between age, size, and environment obscures their independent fitness contributions. We isolated the fitness effects of these factors by experimentally manipulating the correlation between age at flowering and date of flowering in Brassica rapa. We staggered the planting dates of families with differing ages at flowering to produce experimental populations in which age at flowering and date of flowering were positively, negatively, or uncorrelated. In all populations, plants with an early date of flowering produced more seed than those flowering late, regardless of age or size at flowering onset. The temporal environment was thus the principal driver of selection on flowering time, but its importance relative to that of age and size varied with the presence/absence of herbivores and seed predators.     

Brood desertion in Kentish plover: the value of parental care

To understand the evolution of parental care, one needs to estimatethe payoffs from providing care for the offspring and from terminatingcare and deserting them. In this study we estimated the payofffrom care provision, and in a companion paper we analyze thepayoff from offspring desertion. In the current study we experimentallyinvestigated the influence of the number and sex of attendingparents on growth and survival of offspring in the Kentish ploverCharadrius alexandrinus, in two sites (A and B). Either the maleor the female parent was removed from some broods at hatchingof the chicks (female-only and male-only broods, respectively),whereas in control broods both parents were allowed to attendtheir young. At site A survival of the chicks was lower in uniparental(male-only and female-only) broods than in control broods, whereaswe found no difference in brood survival at site B. Brood survivaldecreased over the season. Removal of either parent did not influencethe growth of the young, although growth varied over the breeding season,and it was significantly different between the sites. Theseresults suggest that the payoff from parental care decreasesover the breeding season and that the value of parental care(i.e., the contribution of parents to the survival of theiryoung) may depend on the environment.     

Prediction of Arctic plant phenological sensitivity to climate change from historical records

The pace of climate change in the Arctic is dramatic, with temperatures rising at a rate double the global average. The timing of flowering and fruiting (phenology) is often temperature dependent and tends to advance as the climate warms. Herbarium specimens, photographs, and field observations can provide historical phenology records and have been used, on a localised scale, to predict species’ phenological sensitivity to climate change. Conducting similar localised studies in the Canadian Arctic, however, poses a challenge where the collection of herbarium specimens, photographs, and field observations have been temporally and spatially sporadic. We used flowering and seed dispersal times of 23 Arctic species from herbarium specimens, photographs, and field observations collected from across the 2.1 million km² area of Nunavut, Canada, to determine (1) which monthly temperatures influence flowering and seed dispersal times; (2) species’ phenological sensitivity to temperature; and (3) whether flowering or seed dispersal times have advanced over the past 120 years. We tested this at different spatial scales and compared the sensitivity in different regions of Nunavut. Broadly speaking, this research serves as a proof of concept to assess whether phenology–climate change studies using historic data can be conducted at large spatial scales. Flowering times and seed dispersal time were most strongly correlated with June and July temperatures, respectively. Seed dispersal times have advanced at double the rate of flowering times over the past 120 years, reflecting greater late‐summer temperature rises in Nunavut. There is great diversity in the flowering time sensitivity to temperature of Arctic plant species, suggesting climate change implications for Arctic ecological communities, including altered community composition, competition, and pollinator interactions. Intraspecific temperature sensitivity and warming trends varied markedly across Nunavut and could result in greater changes in some parts of Nunavut than in others.